Cyclostome Bryozoans from the Zalas Quarry, Southern Poland

Callovian (Middle Jurassic) cyclostome bryozoans from the Zalas Quarry, southern Poland

MICHA£ ZATOÑ, URSZULA HARA, PAUL D. TAYLOR & MICHA£ KROBICKI

The specimen-rich and diverse bryozoan fauna from the Callovian hardground at Zalas Quarry in southern Poland is described. Twenty-two taxa of cyclostomes are recorded, of which three species are proposed as new: Microeciella calloviana, Reptomultisparsa viskovae and Mesonopora walteri. Due to preservational problems and an insufficient number of fertile colonies, species-level determination of fourteen forms was not possible. The most common bryozoans present are sheet-like bereniciform colonies, with uniserial runners and oligoserial ribbons less abundant. The number of the Callovian taxa present in the Zalas Quarry is very similar to the Upper Bathonian–Lower Callovian bryozoan assemblage from the classic locality of Balin in southern Poland. Taking the strictly Callovian age into account, the Zalas assemblage is the most diverse for that age ever noted. • Key words: Bryozoa, Cyclostomata, Callovian, Jurassic, Poland.

ZATOŃ, M., HARA, U., TAYLOR, P.D. & KROBICKI, M. 2013. Callovian (Middle Jurassic) cyclostome bryozoans from the Zalas Quarry, southern Poland. Bulletin of Geosciences 88(4), 837–863 (19 figures). Czech Geological Survey, Prague. ISSN 1214-1119. Manuscript received July 8, 2013; accepted in revised form September 5, 2013; published on- line October 21, 2013; issued October 31, 2013.

Michał Zatoń (corresponding author), University of Silesia, Faculty of Earth Sciences, Będzińska Street 60, PL-41-200 Sosnowiec, Poland; [email protected] • Urszula Hara, Polish Geological Institute – National Research Institute, Rakowiecka 4, PL-00-975 Warszawa, Poland; [email protected] • Paul D. Taylor, Natural History Museum, De- partment of Earth Sciences, Cromwell Road, London SW7 5BD, United Kingdom; [email protected] • Michał Krobicki, Polish Geological Institute – National Research Institute; Upper Silesian Branch, Królowej Jadwigi 1, PL-41-200 Sosnowiec, Poland, & AGH University of Science & Technology, Department of General Geology and Geotourism, al. Mickiewicza 30, PL-30-059 Kraków, Poland; [email protected], [email protected]

The Middle Jurassic was a significant time in the evolutio- material described by Reuss from Balin was recently revised nary radiation of the cyclostome bryozoans, which reached by Taylor (2009), who described a total of twenty-three taxa, their maximum diversity for the Jurassic in the Bathonian among which eight were considered to be new. This study (Taylor & Larwood 1990, Jablonski et al. 1997, Taylor & emphasized the morphology of both the gonozooids and Ernst 2008). This peak is due mainly to the exceptional di- pseudopores as important taxonomic characters. Other stud- versity of cyclostomes in Calvados, northern France, parti- ies undertaken in the past few years on the Middle Jurassic cularly the sponge reef and hardground settings of the Cail- (Late Bajocian and Bathonian) cyclostomes of southern Po- lasse de la Basse Ecarde at St Aubin-sur-Mer, where 33 land have revealed the presence of twenty-nine species be- species were recorded by Walter (1970). Research on cyc- longing to nine genera (Zatoń & Taylor 2009, 2010). A few lostome bryozoans from other sites is needed to test whet- recent publications on younger faunas of cyclostomes from her the Bathonian diversity peak is a real phenomenon, or the Polish Upper Jurassic have enhanced our understanding merely reflects the exposure of suitable facies. Although of the systematics and distributional patterns of these bryo- published data on Jurassic cyclostome bryozoans is still zoans (see Hara 2007; Hara & Taylor 1996, 2009). scarce, recent progress towards understanding their diver- The current paper provides the first detailed systematic sity has been made in both North America (see Taylor & description of the Callovian cyclostome bryozoans from Wilson 1999) and central and eastern Europe (see Hara & the Zalas Quarry near Kraków in the southern Poland, de- Taylor 1996, 2009; Taylor 2009; Viskova 2006, 2007, scribing twenty-two taxa belonging to eleven genera. 2008, 2009; Zatoń & Taylor 2009, 2010). Palaeoecological factors are discussed (cf. Hara 2007), The cyclostome bryozoans of the Upper Bathon- such as the substrate type and colony growth-forms of the ian–Lower Callovian of southern Poland were first system- bryozoans that colonized hard, mostly organic shelly sub- atically studied in the 19th century by Reuss (see Reuss strates, often represented by shells of the limid bivalve 1867) from the deposits at the classic locality of Balin. The Ctenostreon proboscideum (Sowerby).

DOI 10.3140/bull.geosci.1466 837 Bulletin of Geosciences Vol. 88, 4, 2013

As the Balin cyclostomes revised by Taylor (2009) Well-preserved uniserial (stomatoporids) and oligoserial come from condensed deposits of Late Bathonian–Early (Oncousoecia) colonies, as well as all bereniciform colo- Callovian and probably also earliest Middle Callovian age nies that were fertile (i.e., possessed gonozooids), were (Mangold et al. 1996), the new fauna described here not sawn off the shell substrate, cleaned ultrasonically, and only supplements this data, but also significantly enlarges examined in an uncoated state using a LEO 1455VP low our knowledge about cyclostome bryozoans of strictly vacuum scanning electron microscope housed at the Natu- Callovian age. ral History Museum in London (NHMUK), or a Philips XL 30 environmental scanning electron microscope housed at the Faculty of Earth Sciences in Sosnowiec, Poland. Ima- Material and methods ges were generated using back-scattered electrons (BSE detector). Material and its provenance. – The Callovian cyclostome It must be stressed that although ultrasonically cleaned, bryozoans were found encrusting various types of fossils many colonies were still coated by a ferruginous crust that (bivalve shells, belemnite rostra, cephalopod moulds) res- obscured some morphological details. Some bereniciform ting on the hardground surface located in the Zalas Quarry, colonies additionally had eroded gonozooids, which al- situated about 8 km south of Krzeszowice and 30 km west lowed generic identification but hampered species-level of Kraków in southern Poland (Fig. 1A, B). The hard- determination due to the lack of key features such as the ground developed on the top of sandy crinoidal limestones position, size and shape of the ooeciopore. of Lower Callovian age (koenigi and calloviense zones) In the present paper the term gonozooid is used for the during a period of non-deposition and associated chemical entire fertile zooid whereas brood chamber refers to its dis- and physical erosion. Features of the hardground are well tal, dilated part with a densely pseudoporous frontal wall in expressed in the presence of limonitic coatings and abun- which the embryos were actually brooded. dant encrusters and borers (Giżejewska & Wieczorek The specimens are housed at the Faculty of Earth Sci- 1976, Dembicz & Praszkier 2007, Zatoń et al. 2011). It is ences, Sosnowiec, with registration numbers prefaced by considered that the period of non-deposition during hard- GIUS 8-3589. ground formation spanned the Middle Callovian (jason and nearly all of the coronatum chrons, see Giżejewska & Wieczorek 1976) and part of the Late Callovian (Matyja Systematic palaeontology 2006, Dembicz & Praszkier 2007). Depressions in the top of the crinoidal limestone are also filled with a nodular de- Order Cyclostomata Busk, 1852 posit (a limestone with nodular structure, termed nodular Family Stomatoporidae Pergens & Meunier, 1886 limestones by Giżejewska & Wieczorek (1976), consisting of clasts and fossils with ferruginous coatings and encrus- Genus Stomatopora Bronn, 1825 tations, derived from the underlying crinoidal limestones. Their chemical corrosion and erosional truncation also in- Type species.–Alecto dichotoma Lamouroux, 1821, by dicate that they were redeposited (Giżejewska & Wieczo- monotypy. rek 1976, Dembicz & Praszkier 2007). The hardground and nodular limestones are overlain by an uppermost Callovian Stomatopora cf. dichotomoides (d’Orbigny, 1850) (lamberti Zone) stromatolitic layer (Giżejewska & Wiec- Figure 2A–C zorek 1976, Matyja 2006, Dembicz & Praszkier 2007). The entire Callovian section is a deepening-upwards sequence Material. – Many variably preserved colonies on several from clastic and carbonate-clastic to strictly pelagic carbo- bivalve shells (GIUS 8-3589). One colony, GIUS nates, and most probably originated in an open outer shelf 8-3589-Z10/1was selected for SEM. environment (cf. Matyja 2006). Although a variety of fossils are encrusted, most of the Description. – Colony encrusting, uniserial, the single bryozoans encrust shells of the large limid bivalve Cteno- scanned example consisting of five zooids forming a single streon proboscideum (Sowerby) (Fig. 1C, D), which also branch, not ramifying (Fig. 2A); branch curved near to the hosted other diverse and abundant encrusters and borings base of the third zooid, the three youngest zooids growing (see Zatoń et al. 2011 for details). The encrusted bivalves alongside a serpulid tube. Ancestrula long, about 400 μm were collected from the uppermost part of the crinoidal (including protoecium), bending by about 140° at mid- limestones – hardground surface. length. Protoecium a low dome, broad compared to proximal part of ancestrular tube, about 130 μm in transverse Methods. – Each encrusted fossil was inspected under a bi- diameter, subcircular in outline, the proximal edge contain- nocular microscope and every bryozoan colony marked. ing a few embayments; surface well preserved, smooth

838 Micha³ Zatoñ et al. Callovian (Middle Jurassic) cyclostome bryozoans from southern Poland

A B

1 cm C 1 cm D

Figure 1. Locality, section sampled and material.•A–geological sketch-map of the environs of Krzeszowice with location of Zalas Quarry indicated by an asterisk (from Zatoń et al. 2011 and simplified after Nawrocki et al. 2007); 1 – Cenozoic mudstones, 2 – Mesozoic deposits, 3 – Permian rhyodacite domes and rhyolitic, trachytic and andesitic dykes, 4 – Permo-Carboniferous basaltic lavas and trachyandesite sills, 5 – Upper Carboniferous conglomer- ates, 6 – Lower Carboniferous mudstones.•B–Middle Jurassic section exposed at the Zalas Quarry, 1 – Lower Callovian crinoidal limestones, 2 – hardground (sampled interval) of presumably Middle-Late Callovian age, 3 – bed with stromatolites and oncolites of latest Callovian age, 4 – Lower Oxfordian limestones. • C, D – sampled Ctenostreon bivalve shells showing intensive encrustation and boring by various sclerobionts. Cyclostome bryozoans colonies are indicated by arrows. Scale bars equal 1 cm. without any obvious pseudopores but pierced by an irre- of this Zalas form match those of S. dichotomoides as inter- gular hole 20 μm in diameter, possibly a boring (Fig. 2B). preted by Hara & Taylor (2009) from the Polish Lower Ancestrular tube initially 60 μm in width, enlarging to Kimmeridgian, the pseudopores are much sparser. Unfor- 80 μm distally, tube marked by transverse growth checks; tunately, no information is available about these structures aperture broken, about 60 μm in transverse diameter. in type or topotype material of this species from the French Budded autozooids with apertures about 80 μm in trans- Bajocian. verse diameter, the youngest zooid measuring 800 μm long by about 200 μm wide. Pseudopores widely scattered, apparently lacking from some parts of the frontal Stomatopora cf. recurva Waagen, 1867 wall, teardrop-shaped, pointed distally (Fig. 2C), about Figure 2D–F 7 μm long by 4 μm wide. Material. – Many variably preserved colonies on bivalve Remarks. – For an abbreviated synonymy of S. dichotomoi- shells. Three colonies were selected for SEM: GIUS des, see Hara & Taylor (2009). While the zooid dimensions 8-3589-Z7/4, GIUS 8-3589-Z7/5 and GIUS 8-3589-Z3b.

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Figure 2. Stomatoporid cyclostome bryozoans from the Callovian hardground of the Zalas Quarry, southern Poland. • A–C – Stomatopora cf. dichotomoides (d’Orbigny, 1850), GIUS 8-3589-Z10/1. A – initial, uniserial part of the colony; B – ancestrula with disc-shaped protoecium and curved distal tube; C – autozooid frontal wall with scattered pseudopores. • D–F – Stomatopora cf. recurva Waagen, 1867, GIUS 8-3589-Z7/4. D – early astogeny; E – ancestrula with pseudoporous protoecium; F – dichotomous bifurcation showing autozooid frontal walls.

Description. – Colony encrusting, uniserial/pseudounise- dome, 360 μm in transverse diameter, with longitudinally rial, branches bifurcating every second or third zooid, the elliptical, moderately abundant pseudopores scattered two daughter zooids remaining linked for up to a third of over the smooth surface (Fig. 2E). Budded autozooids their lengths after bifurcation (Fig. 2D). First bifurcation large, stout, about 800 μm long by 600 μm maximum at ca 120°, second at 90°, later bifurcations ranging down width (branch width), with subcircular or transversely el- to ca 60°. Ancestrula short, straight, stout, 550 μm long liptical apertures 100 μm long by 100–130 μm wide (including protoecium), up to 280 μm wide; aperture sub- (Fig. 2F). Pseudopores subcircular, ca 10 μm in diameter circular, transverse diameter 90 μm. Protoecium a low (Fig. 2E).

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Figure 3. Stomatopora sp. from the Callovian hardground of the Zalas Quarry, southern Poland, GIUS 8-3589-Z3/9.•A–colony associated with serpulid tubes; B – ancestrula with narrow protoecium; C – autozooid frontal walls; D – pseudopores.

Remarks. – This taxon is probably pseudouniserial, as decreasing from an initial 180° to less than 90° by the third indicated by the comparatively large width of the bran- generation of branching (Fig. 3A). Ancestrula short, about ches and change in slope along the branch edges, imply- 370 μm long (including protoecium), straight, up to 170 μm ing that the median zooid is flanked by the proximal in maximum width; aperture subcircular, 80 μm in diame- parts of the next distal zooid in series, as well as the dela- ter. Ancestrular tube almost as wide as protoecium. Protoe- yed separation of zooids after each bifurcation. The cium comparatively flat, about 180 μm in transverse dia- large size of the zooids and width of the branches allows meter, the surface rugose (Fig. 3B), facetted; pseudopores the form to be compared with S. recurva Waagen, which irregularly scattered, about 5 μm in diameter, subcircular to was recorded from the nearby locality of Balin by Taylor teardrop-shaped and pointed distally. Budded autozooids (2009). As with the other established species of Stomato- up to 1100 μm long by 440 μm wide, with subcircular aper- pora, however, critical details of the pseudopores and tures 130–140 μm wide (Fig. 3C). Pseudopores regularly early astogeny in the German type material of S. recurva spaced, about 30 μm apart (centre to centre), teardrop- are unknown, hampering the precise characterization of shaped with pointed distal ends, with a sunken, iris-like S. recurva. ring of calcification defining a central opening 2–3 μm in diameter (Fig. 3D).

Stomatopora sp. Remarks. – A third, unnamed taxon of Stomatopora Figure 3A–D from Zalas has autozooids intermediate in size (width) between those of S. cf. dichotomoides and S. cf. recurva. Material. – One colony, GIUS 8-3589-Z3/9. The ancestrula is distinctive in possessing a flat protoecium with sparse pseudopores, while at least some of the Description. – Colony encrusting, uniserial, branches bi- pseudopores on the budded zooids have a distinctive furcating every alternate zooid with angles of bifurcation iris-like constriction.

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Family Oncousoeciidae Canu, 1918 diameter, spaced approximately 20 μm apart (Fig. 4F). Go- nozooid longitudinally ovate (Fig. 4E), brood chamber Genus Oncousoecia Canu, 1918 >600 μm long by 500 μm wide, densely pseudoporous, the spacing between pseudopores roughly equivalent to their Type species.–Tubulipora lobulata Canu, 1918 (see Tay- diameters; ooeciopore longitudinally elliptical, conside- lor & Zatoń 2008). rably smaller than an autozooidal aperture, about 70 μm long by 60 μm wide. Oncousoecia sp. 1 Figure 4A–C Remarks. – This form has appreciably larger apertures than Oncousoecia sp. 1 and the pseudopores are circular rather Material. – Many (ca 34), variably preserved and ?infertile than slit-like. Despite being fertile, the material is inadequ- colonies encrusting several bivalve shells (GIUS 8-3589), ate to allow a reliable comparison with established species of which four were selected for SEM: GIUS 8-3589-30/1, of Oncousoecia, which in any case are poorly characterized GIUS 8-3589-Z5/1, GIUS 8-3589-M3, GIUS in the literature. The species of Oncousoecia from Balin, 8-3589-M15a/1. apart from being represented by infertile colonies, differs in having longitudinally elliptical pseudopores (Taylor 2009). Description. – Colony encrusting, pluriserial, comprising bifurcating, ribbon-like branches, with 2–7 autozooids across the width of the branch; kenozooids developed on Genus Microeciella Taylor & Sequeiros, 1982 sloping lateral edges of at least some branches (Fig. 4A). Ancestrula about 300 μm long with a protoecium 180 μm in Type species.–Microeciella beliensis Taylor & Sequeiros, transverse diameter; surface too poorly preserved in the 1982 by original designation. single example to reveal pseudopore shape or patterning. Autozooids elongate, frontal walls convex, about Microeciella calloviana sp. nov. 500–1000 μm long by 300 μm wide; apertures subcircular Figure 5A–F or longitudinally elliptical, ca 100 μm in diameter; peristome short to moderate in length, distally tapering and inclined at Types. – Holotype: GIUS 8-3589-Z5/2 (Fig. 5A–D); para- 60° or more to colony surface. Pseudopores elongate, slit-like, types: GIUS 8-3589-M6 (Fig. 5E), GIUS 8-3589-M7 about 15 μm long, becoming longitudinally elliptical in weat- (Fig. 5F), GIUS 8-3589-Z3a/01. hered specimens (Fig. 4C). Gonozooid not unequivocally identified, only a longitudinally ovate swelling observed in Type locality. – Zalas Quarry, southern Poland. one specimen being a possible gonozooid (Fig. 4B). Type horizon. – Callovian hardground, Middle Jurassic. Remarks. – The lack of an unequivocal gonozooid precludes species-level identification of this form. However, it is Etymology. – After Callovian, the age of the species descri- not the same species described among the Reuss material bed. from Balin by Taylor (2009), which has significantly larger zooids (up to 1880 μm long by 330–400 μm wide). Material. – Four colonies encrusting bivalve shells: GIUS 8-3589-Z5/2, GIUS 8-3589-M6, GIUS 8-3589-M7, GIUS 8-3589-Z3a/01. Oncousoecia sp. 2 Figure 4D–F Measurements. – Autozooid frontal wall length: 570–670 μm; autozooid frontal wall width: 140–160 μm; longitudinal Material. – One fertile but incomplete colony encrusting a apertural diameter: 87–110 μm; transverse apertural dia- bivalve shell: GIUS 8-3589-Z3/2a. meter: 87–100 μm; gonozooid total length: 1030 μm; brood chamber length: 530–630 μm; gonozooid width: 600–820 μm; Description. – Colony encrusting, pluriserial, comprising, ooeciopore length: 30 μm; ooeciopore width: 50 μm; pseu- ribbon-like branches (presumably bifurcating), 2–3 auto- dopore width: 6–8 μm. zooids wide (Fig. 4D). Ancestrula and early astogenetic stages not observed. Autozooids large, elongate, frontal Diagnosis.–Microeciella species having a sheet-like co- walls convex, about 800–1300 μm long by 300 μm wide, lony possessing elongate autozooids of a narrow width zooecial boundaries well defined by grooves; apertures range and gonozooids with ovoidal brood chamber with subcircular, 140–170 μm in diameter; peristome short to terminally located ooeciopores. Pseudopores circular to moderate in length. Pseudopores circular, about 7 μm in wide teardrop-shaped.

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Figure 4. Oncousoeciid cyclostome bryozoans from the Callovian hardground of the Zalas Quarry, southern Poland. • A–C – Oncousoecia sp. 1, GIUS 8-3589-M15a/1. A – oligoserial colony with kenozooid on sloping lateral edge (arrowed); B – probable gonozooid (arrowed); C – pseudopores. • D–F – Oncousoecia sp. 2, GIUS 8-3589-Z3/2a. D – colony fragment with preserved gonozooid (arrowed); E – brood chamber with subterminal ooeciopore (arrowed); F – pseudopores.

Description. – Colony encrusting, sheet-like, berenici- marked by shallow grooves. Peristomes short, tapering form, multiserial, unilamellar (Fig. 5A). Autozooids elon- distally. Apertures semicircular to longitudinally elongate, gate with convex frontal walls; zooidal boundaries distinct, some closed by terminal diaphragms located a little beneath

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Figure 5. Microeciella calloviana sp. nov. from the Callovian hardground of the Zalas Quarry, southern Poland.•A–incomplete sheet-like colony with three gonozooids; B – gonozooid with a broken brood chamber and terminal ooeciopore (arrowed); C – autozooids; D – pseudopores; E, F – gonozooids with small terminal ooeciopores (arrowed). A–D – holotype (GIUS 8-3589-Z5/2), E, F – paratypes; E – GIUS 8-3589-M6; F – GIUS 8-3589-M7. the rim of the peristome (Fig. 5C). Pseudopores circular to ony; roof densely pseudoporous with edges indented by wide teardrop-shaped (Fig. 5D). neighbouring autozooids. Ooeciopore terminal, semicircu- Gonozooids (Fig. 5B, E, F) common, occurring in two lar, half the size of an autozooidal aperture, situated on a generations in the holotype (Fig. 5A). Proximal frontal short, upright ooeciostome (Fig. 5B, E, F). wall indistinguishable from that of an autozooid. Brood chamber ovoidal in shape, inflated, varying from slightly Remarks. – Gonozooid shape, ooeciopore size and loca- longer than wide to wider than long even in the same col- tion, as well as pseudopore characteristics, are similar in all

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Figure 6. Microeciella aff. annae Zatoń & Taylor, 2009 from the Callovian hardground of the Zalas Quarry, southern Poland, GIUS 8-3589-C/W-230/2.•A–sheet-like colony with four gonozooids; B – autozooids; C – gonozooid with terminal ooeciopore (arrowed); D – close-up of the tiny ooeciopore (centre), note the elongate, slit-like pseudopores on both the autozooidal and brood chamber frontal walls. four colonies included in this species. Some variations in have subterminal ooeciopores (see Hara & Taylor 2009; Tay- brood chamber shape (width/length ratio) are present, even lor 2009; Taylor & Sequeiros 1982; Taylor & Wilson 1999; in the same colony, but brood chamber size is similar in the Zatoń & Taylor 2009, 2010), whereas the ooeciopore is termi- colonies, as are autozooidal frontal wall and aperture sizes, nal in M. calloviana. although some of the autozooids are wider in one paratype (GIUS 8-3589-M7). With respect to the general appearance of the gonozooids Microeciella aff. annae Zatoń & Taylor, 2009 and the terminal position of the ooeciopore, this species is Figure 6A–D reminiscent of the Bathonian species Microeciella annae described by Zatoń & Taylor (2009) from the Polish Jura. How- Material. – One colony encrusting a bivalve shell, GIUS ever, the autozooids are distinctly smaller and pseudopores 8-3589-C/W-230/2. are narrowly teardrop-shaped and spindle-like in the latter species. Another similar Bathonian taxon, M. aff. annae, also Measurements. – Autozooid frontal wall length: 410–550 μm; differs in having smaller autozooids, ovoidal gonozooids that autozooid frontal wall width: 140–170 μm; longitudinal are longer than wide, and transversely elliptical pseudopores apertural diameter: 70–100 μm; transverse apertural dia- (see Zatoń & Taylor 2010). The Zalas species shows some meter: 75–90 μm; gonozooid total length: 860–1120 μm; similarities with Microeciella maleckii from the Upper brood chamber length: 460–650 μm; gonozooid width: Bathonian of the Polish Jura (Zatoń & Taylor 2009). How- 600–790 μm; ooeciopore length: 50 μm; ooeciopore width: ever, apart from the slightly smaller apertures, the latter spe- 50 μm; pseudopore length: 14–18 μm. cies also differs in the subterminal location of the ooeciopore and the more teardrop-shaped pseudopores. In general, spe- Description. – Colony encrusting, sheet-like, berenici- cies of Microeciella, including the type species M. beliensis, form, multiserial, unilamellar (Fig. 6A). Autozooids short

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Figure 7. Microeciella cf. matisconensis Walter, 1970 from the Callovian hardground of the Zalas Quarry, southern Poland.•A–colony showing three gonozooids, close to the colony margin; B – autozooids associated with two gonozooids having subterminal ooeciopores (arrowed); C – another gonozooid with subterminal ooeciopore (arrowed); D – pseudopores. A, B, D – GIUS 8-3589-Z19/1, C – GIUS 8-3589-Z6. with slightly convex frontal walls; zooidal boundaries shal- the Zalas taxon differs in its larger gonozooids and low-grooved. Peristomes short, upright, tapering distally. slit-like pseudopores. Gonozooid shape and pseudopore Apertures circular to semicircular (Fig. 6B). Pseudopores characteristics are similar to Microeciella sp. from the dense, long, slit-like (Fig. 6D). Upper Bathonian–Lower Callovian of Balin (see Taylor Gonozooids common, occurring in two generations, 2009), although the ooeciopore is subterminal in the Balin best developed at the colony margins (Fig. 6A). Proximal species. part indistinguishable from an autozooidal frontal wall. Brood chamber densely pseudoporous, inflated, ovoidal in shape, slightly wider than longer or equidimensional, with Microeciella cf. matisconensis (Walter, 1970) margins indented by neighbouring autozooids (Fig. 6C). Figure 7A–D Ooeciopore terminal, circular, smaller than an autozooid aperture, situated on a short, upright ooeciostome Material. – Four colonies encrusting bivalve shells: GIUS (Fig. 6D). 8-3589-Z5/2, GIUS 8-3589-Z6, GIUS 8-3589-Z19/1, GIUS 8-3589-M12/2. Remarks. – With respect to autozooid length and gonozooid shape, this taxon is most similar to Microeciella an- Measurements. – Autozooid frontal wall length: 470–800 μm; nae from the Bathonian of the Polish Jura (Zatoń & Tay- autozooid frontal wall width: 100–170 μm; longitudinal lor 2009). The latter species, however, is characterised by apertural diameter: 70–100 μm; transverse apertural dia- slightly smaller apertures, an ooeciopore twice the size of meter: 60–90 μm; gonozooid total length: 720–970 μm; the present species, and more teardrop to spindle-like brood chamber length: 320–470 μm; gonozooid width: pseudopores. From another similar Lower Bathonian 290–430 μm; ooeciopore length: 50 μm; ooeciopore width: form, assigned to M. aff. annae by Zatoń & Taylor (2010), 50 μm; pseudopore length: 12–15 μm.

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200 µm C 20 µm D

Figure 8. Microeciella cf. maleckii Zatoń & Taylor, 2009 from the Callovian hardground of the Zalas Quarry, southern Poland, GIUS 8-3589-M8. • A – part of a sheet-like colony; B – autozooids; C – gonozooid with subterminal ooeciopore (white arrow), note the neighbouring autozooid embraced by the brood chamber (black arrow); D – pseudopores.

Description. – Colony encrusting, sheet-like, berenici- M. matisconensis, making direct comparison difficult and form, multiserial, unilamellar (Fig. 7A). Autozooids leaving some doubt about the conspecificity of the French long, with slightly convex frontal walls; zooidal bounda- and Polish material. The Microeciella sp. of Taylor ries variably distinct, shallowly grooved. Peristomes (2009) from the Upper Bathonian–Lower Callovian of short, tapering distally. Apertures circular to slightly Balin, southern Poland, is similar with respect to the sub- longitudinally elongate (Fig. 7B). Pseudopores long, terminal position of the ooeciopore and slit-like pseudo- slit-like (7D). pores; however, the brood chambers are larger and wider, Gonozooids common, occurring in up to two genera- and the ooeciopore transversely elongated in this species. tions (Fig. 7A). Proximal frontal wall indistinguishable The Zalas form is similar to other Jurassic species of Mic- from that of an autozooid. Brood chambers small, in- roeciella in the subterminal position of the ooeciopore but flated, globular in shape, slightly longer than wide; edges differs in having characteristically small, globular brood indented by neighbouring autozooids. Ooeciopore circu- chambers. lar in outline, subterminal, smaller than an autozooid aperture (Fig. 7B, C). Microeciella cf. maleckii Zatoń & Taylor, 2009 Remarks. – Autozooid width and gonozooidal characte- Figure 8A–D ristics in the Zalas material are identical to those of the French species Microeciella matisconensis (Walter, Material. – One colony encrusting bivalve shell, GIUS 1970). However, pseudopore morphology is unknown in 8-3589-M8.

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Measurements. – Autozooid frontal wall length: 380–400 μm; Type horizon. – Callovian hardground, Middle Jurassic. autozooid frontal wall width: 100–120 μm; longitudinal apertural diameter: 70–90 μm; transverse apertural diame- Etymology. – In honor of Lena A. Viskova (PIN, Moscow), ter: 50–60 μm; gonozooid total length: 620 μm; brood in recognition of her extensive contributions to the study of chamber length: 600 μm; gonozooid width: 550 μm; ooecio- post-Palaeozoic bryozoans. pore length: 50–60 μm; ooeciopore width: 40 μm; pseudopore length: 8–10 μm; pseudopore width: 6–7 μm. Material. – One colony encrusting a bivalve shell, GIUS 8-3589-Z5/1 (holotype). Description. – Colony encrusting, sheet-like, bereniciform, multiserial, unilamellar (Fig. 8A). Autozooids elon- Measurements. – Autozooid frontal wall length: 530–600 μm; gate with slightly convex frontal walls; zooidal bounda- autozooid frontal wall width: 110–140 μm; longitudinal ries distinct distally, shallowly grooved. Peristomes short, apertural diameter: 60–90 μm; transverse apertural diame- tapering distally. Apertures semicircular to oval ter: 60–90 μm; gonozooid total length: 1040 μm; brood (Fig. 8B). Pseudopores teardrop-shaped to oval in outline chamber length: 640 μm; gonozooid width: 390 μm; ooeci- (Fig. 8D). opore length: 70 μm; ooeciopore width: 90 μm; pseudopore Gonozooids infrequent, the preserved example hav- length: 8–10 μm; pseudopore width: 12–15 μm; ancestrula ing a proximal frontal wall indistinguishable from an length: 190 μm; protoecium length: 140 μm; protoecium autozooid frontal wall. Brood chamber ovoidal, with width: 130 μm. convex frontal wall, widening distally, its margin indented by neighbouring autozooids; right distal end em- Diagnosis. – Reptomultisparsa species having elongate au- bracing one of the autozooidal peristomes and then re- tozooids, spindle-shaped brood chamber with subterminal joining the rest of the brood chamber close to the ooeciopore, and transversely elliptical pseudopores partly ooeciopore (Fig. 8C). Ooeciopore circular, smaller than closed by sunken walls at their proximal and distal ends an autozooid aperture, situated on a short, subterminal leaving a transverse, slit-like opening. ooeciostome. Description. – Colony encrusting, fan-shaped, multiserial, Remarks. – The ovoidal, distally widening brood cham- unilamellar, small (Fig. 9A). Ancestrula with circular, con- ber with subterminal ooeciopore and marginal indenta- vex, smooth protoecium; distal tube straight, short; one dis- tion by the neighbouring autozooids invite comparison tal autozooid budded from ancestrula. between this form and the Upper Bathonian species Autozooids elongate with slightly convex frontal walls; M. maleckii Zatoń & Taylor, 2009. However, the com- zooidal boundaries well marked by shallow grooves plete envelopment of an autozooidal peristome by the (Fig. 9B, C). Peristomes short, tapering distally. Apertures brood chamber evident in the colony from Zalas is un- semicircular to circular (Fig. 9C). Pseudopores transver- known in M. maleckii and other Jurassic species of Mic- sely elliptical, partly closed by sunken walls at their proxi- roeciella. With respect to other morphological features mal and distal ends leaving a transverse, slit-like opening of the autozooids, brood chamber and pseudopores, the (Fig. 9D). two species are similar to each other, M. maleckii diffe- Only one gonozooid preserved (Fig. 9A, B); its proxi- ring from the Zalas form only in the slightly longer auto- mal frontal wall long, convex, indistinguishable from an zooidal frontal walls. autozooidal frontal wall. Brood chamber longer than wide, spindle-shaped, convex, with edges indented by apertures of neighbouring autozooids at mid-length. Roof densely Family Multisparsidae Bassler, 1935 pseudoporous. Ooeciopore subterminal, semi-circular, smaller than an autozooidal aperture, situated on a short, Genus Reptomultisparsa d’Orbigny, 1853 upright ooeciostome (Fig. 9C).

Type species.–Diastopora incrustans d’Orbigny, 1850 Remarks. – From other representatives of the genus Repto- (Taylor 1984). multisparsa, this new species differs in its transversely elliptical pseudopores partly closed by sunken walls along their Reptomultisparsa viskovae sp. nov. proximal and distal sides. Although the material consists of Figure 9A–D only one colony, these unusual pseudopores allow the new species to be established. Partial occlusion of pseudopores Types. – Holotype: GIUS 8-3589-Z5/1 (Fig. 9A–D). by ring-like walls or spines has been described previously in cyclostome bryozoans (e.g., Weedon & Taylor 1997) but the Type locality. – Zalas Quarry, southern Poland. mode of closure in R. viskovae appears to be unique.

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Figure 9. Reptomultisparsa viskovae sp. nov. from the Callovian hardground of the Zalas Quarry, southern Poland, GIUS 8-3589-Z5/1.•A–small colony overgrowing another cyclostome, with ancestrula (arrowed); B – autozooids and gonozooid having subterminal ooeciopore (arrowed); C – close-up of the colony growing edge; D – pseudopores, note microendoliths in lower right.

Reptomultisparsa aff. kawodrzanensis Zatoń & Taylor, Gonozooids common, with proximal frontal wall simi- 2010 lar in width to an autozooid frontal wall. Brood chamber Figure 10A–D longitudinally elongated, much longer than wide, ovoidal in shape; edges indented by neighbouring autozooids. Material. – One colony encrusting a bivalve shell, GIUS Ooeciopore subterminal, subcircular, slightly wider than 8-3589-Z2. longer, smaller than an autozooidal aperture (Fig. 10C).

Measurements. – Autozooid frontal wall length: 420–800 μm; Remarks. – With respect to autozooid shape and size, go- autozooid frontal wall width: 125–175 μm; longitudinal aper- nozooid morphology and pseudopore characteristics, the tural diameter: 110–140 μm; transverse apertural diameter: form described here is most similar to Reptomultisparsa 87–120 μm; gonozooid total length: 1280–1450 μm; brood kawodrzanensis Zatoń & Taylor, 2010 from the Lower chamber length: 720–1100 μm; gonozooid width: 320 to Bathonian of the Polish Jura. It differs, however, in hav- 420 μm; ooeciopore length: 90 μm; ooeciopore width: 110 μm; ing a generally shorter brood chamber (720–110 μm pseudopore length: 6–8 μm; pseudopore width: 6–8 μm. vs. 940–1720 μm) and lacking a distinct ooeciostome, although the latter may be a preservational artefact. Description. – Colony encrusting, sheet-like, bereniciform, Another species, Reptomultisparsa harae Zatoń & Tay- multiserial, unilamellar (Fig. 10A). Autozooids elongate with lor, 2009 from the Upper Bajocian and Middle Batho- convex frontal walls; zooidal boundaries well marked with nian of England and Poland, has a similar gonozooid but shallow grooves. Peristomes short, tapering distally. Apertu- differs in its long, slit-like pseudopores. Despite these res semicircular to longitudinally elongated (Fig. 10B). Pseu- differences from existing species of Reptomultisparsa, dopores circular to teardrop-shaped, unclear due to the worn more material is needed to warrant the establishment of a colony surface (Fig. 10D). new species.

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Figure 10. Reptomultisparsa aff. kawodrzanensis Zatoń & Taylor, 2010 from the Callovian hardground of the Zalas Quarry, southern Poland, GIUS 8-3589-Z2.•A–fertile, sheet-like colony; B – autozooids; C – complete gonozooid with subterminal ooeciopore (white arrow) and an aborted gonozooid (black arrow); D – pseudopores.

Reptomultisparsa sp. edges indented by apertures of neighbouring autozooids Figure 11A–D for three-quarters of the length (Fig. 11C). Ooeciopore subterminal, semicircular, slightly wider than longer, similar Material. – Two colonies encrusting bivalve shells: GIUS to an autozoidal aperture in width, situated on a very short, 8-3589-M5/3 and GIUS 8-3589-C/W-230/1, the first one upright ooeciostome. being fertile. Remarks. – This taxon is most similar to the Reptomulti- Measurements. – Autozooid frontal wall length: 290–710 μm; sparsa sp. of Hara & Taylor (2009) from the Lower Kim- autozooid frontal wall width: 140–190 μm; longitudinal meridgian of Wierzbica and Małogoszcz in the Holy Cross apertural diameter: 70–130 μm; transverse apertural dia- Mountains, Poland. The latter, however, has a wider and meter: 90–110 μm; brood chamber length: 760 μm; gono- flatter brood chamber crossed by transverse growth bands. zooid width: 650 μm; ooeciopore length: 65 μm; ooecio- It is also reminiscent of Reptomultisparsa tumida Taylor, pore width: 90 μm; pseudopore length: 4 μm; pseudopore 1980 from the Upper Bathonian of England but this slightly width: 4 μm. older species has a more oval brood chamber.

Description. – Colony encrusting, muliserial, unilamellar, bereniciform. Autozooids short, with flat frontal walls; zo- Family Plagioeciidae Canu, 1918 oidal boundaries indistinct (Fig. 11A, B). Peristomes short, tapering distally. Apertures circular (Fig. 11B). Pseudopo- Genus Hyporosopora Canu & Bassler, 1929 res dense, tiny, semicircular to slightly ovoidal in outline (Fig. 11D). Type species. – Hyporosopora typica Canu & Bassler, Gonozooid with broad oval, inflated brood chamber, its 1929, by original designation.

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Figure 11. Reptomultisparsa sp. from the Callovian hardground of the Zalas Quarry, southern Poland, GIUS 8-3589-M5/3.•A–nearly flat colony surface; B – autozooids; C – gonozooid with subterminal ooeciopore (arrowed); D – pseudopores.

Hyporosopora tenera (Reuss, 1867) width: 60–80 μm; pseudopore length: 9–17 μm; pseudo- Figure 12A–F pore width: 2–3 μm.

1867 Berenicea tenera, Reuss, p. 8 (partim), pl. 1, fig. 9. Description. – Colony encrusting, multiserial, berenici- 2009 Hyporosopora tenera (Reuss). – Taylor, p. 37, form, unilamellar, usually a small disc up to 3200 μm in figs 7C, 9A–F. diameter, often elliptical with a maximum width of 2009 Hyporosopora tenera (Reuss). – Zatoń & Taylor, 6400 μm, but also may form small flabelliform colonies; p. 282, fig. 12. growing edge thin, composed of one generation of zooids (Fig. 12A). Material. – Nine colonies encrusting bivalve shells, seven Autozooids small, longitudinally elongate, frontal of which were SEMed: GIUS 8-3589-M12/1, GIUS walls convex, with distinct boundaries, covered by trans- 8-3589-M10, GIUS 8-3589-M2, GIUS 8-3589-230-7, verse ridges spaced irregularly about 18–25 μm apart, in GIUS 8-3589-Z3/8, GIUS 8-3589-Z6a, GIUS-3589-Z1. some places discontinuous, low in profile, present over brood chambers. Apertures oval in shape, sometimes Measurements. – Autozooid frontal wall length: 230–430 μm; nearly circular, arranged roughly in quincunx, more autozooid frontal wall width: 90–130 μm; longitudinal crowded at the colony margin; terminal diaphragms pres- apertural diameter: 60–100 μm; transverse apertural dia- ent but rare, located a little beneath the frontal wall, most of meter: 60–80 μm; gonozooid total length: 570–1100 μm; the apertures sealed by the sediment. Peristomes very nar- brood chamber length: 600 μm; gonozooid width: row, tapering distally in the central part of the colony 450–600 μm; ooeciopore length: 50–60 μm; ooeciopore (Fig. 12B). Pseudopores slit-like, rather long, in some

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100 µm C 100 µm D

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Figure 12. Hyporosopora tenera (Reuss, 1867) from the Callovian hardground of the Zalas Quarry, southern Poland.•A–disc-shaped colony view; B – autozooids; C – gonozooid with subterminal ooeciopore (arrowed); D – pseudopores; E, F – gonozooids from two colonies with subterminal ooeciopore well-visible in F (arrowed). A–D – GIUS 8-3589-Z3/8; E – GIUS 8-3589-230-7; F – GIUS 8-3589-M12/1. cases dagger-shaped, covering the entire colony surface (Fig. 12C, E, F). Ooeciopore terminal, transversely ellipti- (Fig. 12D). cal, smaller that an autozooidal aperture (Fig. 12F). Gonozooids transversely elongate in outline, almost half longer than wide, usually one to few gonozooids pres- Remarks. – This Zalas bryozoan agrees in almost all mor- ent close to the colony margin, distal edge often straight, phological and morphometrical features well with the type margins surrounded densely by the autozooidal apertures. material of H. tenera from the Upper Bathonian–Early Cal- Brood chamber inflated, roughly globular, wider than long lovian of Balin recently revised by Taylor (2009). The main

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Figure 13. Hyporosopora aff. sauvagei (Gregory, 1896) from the Callovian hardground of the Zalas Quarry, southern Poland, GIUS 8-3589-M13. • A – part of a large colony imaged in SE mode; B – autozooids; C – gonozooid with subterminal ooeciopore (arrowed); D – round-shaped pseudopores. differences are the shorter autozooecial frontal walls and Measurements. – Autozooid frontal wall length: 580–770 μm; brood chambers in the Zalas material. Diaphragms, which autozooid frontal wall width: 260–290 μm; longitudinal are common in the Balin Oolite material, are less frequent in apertural diameter: 140–170 μm; transverse apertural dia- the Zalas specimens, and fine growth lines on the colony meter: 120–150 μm; gonozooid total length: 2400 μm; brood surface are better defined in specimens from Balin. chamber length: 2100 μm; gonozooid width: 1500 μm; ooe- Recently described material from the Middle Jurassic ciopore length: 120–130 μm; ooeciopore width: 110 μm; ore-bearing clays of the Polish Jura resembles the specimens pseudopore length: 10–13 μm; pseudopore width: 10–11 μm. from Zalas but the gonozooids are larger (see Zatoń & Taylor 2009, p. 282, fig. 12C). The majority of colonies of H. tenera Description. – Colony large (more than 20 mm in diameter from Zalas are subcircular in outline but others are flabellate. when complete), encrusting, sheet-like, bereniciform, uni- The characteristic longitudinally elongate, narrow, slit-like lamellar (Fig. 13A). pseudopores that densely cover the surface are obscured by Autozooids large, gently convex and slightly sinuous, encrustations of ferrous oxide in some of the specimens. expanded a little below the apertures with well-defined boundaries (Fig. 13B). Peristomes short and narrow, slightly tapering distally. Apertures circular to elliptical, Hyporosopora aff. sauvagei (Gregory, 1896) some closed by terminal diaphragms (Fig. 13A). Pseu- Figure 13A–D dopores very distinctive, wide teardrop-shaped to circular, densely and regularly spaced (Fig. 13D). Material. – One, large incomplete colony encrusting a be- Gonozooids large, brood chamber much broader lemnite guard, GIUS 8-3589-M13. than long, boomerang-shaped with two prominent and

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Figure 14. Hyporosopora cf. enstonensis Pitt & Thomas, 1969 from the Callovian hardground of the Zalas Quarry, southern Poland.•A–growing margin of the fertile colony associated with serpulid tubes; B – autozooids; C – gonozooid with subterminal ooeciopore (arrowed); D – pseudopores. A, C – GIUS 8-3589-Z11; B, D – GIUS 8-3589-Z3. symmetrically arranged distal lobes (Fig. 13A, C). Brood sic species of Hyporosopora, such as Hyporosopora sp. of chamber roof indented at its edges by numerous autozooidal Taylor (1981) from the Tithonian Portland Stone of Dorset, apertures. Ooeciopore terminal, subcircular, slightly longer and H. radomensis Hara & Taylor, 2009 from the Lower then wide, a little smaller than an autozooidal aperture, en- Kimmeridgian of the Holy Cross Mountains, Poland. The circled by a narrow, short ooeciostome. latter differs from the Zalas species in having narrower autozooidal frontal wall widths as well as a smaller Remarks. – The holotype of Hyporosopora sauvagei ooeciopore. (NHM B194) was described by Gregory (1896, p. 82, pl. 3, fig. 4) from the Upper Bathonian Bradford Clay of Bradford-on-Avon, Wiltshire, England. In the overall mor- Hyporosopora cf. enstonensis Pitt & Thomas, 1969 phology of the colony, autozooids and gonozooids, Figure 14A–D Gregory’s species resembles the Callovian specimen from Zalas. However, pseudopores are wide teardrop-shaped to Material. – Two colonies encrusting bivalve shells: GIUS circular in Polish material identified as Hyporosopora aff. 8-3589-Z3/11, GIUS 8-3589-Z11/2. sauvagei (Taylor 2009, Zatoń & Taylor 2009), as well as in the Zalas material studied here, whereas they are Measurements. – Autozooid frontal wall length: 300–430 μm; crescent-shaped in the holotype of H. sauvagei (see Hara & autozooid frontal wall width: 140–180 μm; longitudinal aper- Taylor 2009). In view of the emerging importance of pseu- tural diameter: 100–130 μm; transverse apertural diameter: dopore morphology in cyclostome taxonomy, this diffe- 70–80 μm; gonozooid total length: 750–900 μm; brood cham- rence should be regarded as significant at species level. ber length: 600 μm; gonozooid width: 500–660 μm; ooecio- The distinctive boomerang-shaped gonozooid in the pore length: 40–50 μm; ooeciopore width: 50 μm; pseudopore Zalas specimen studied here also recalls some other Juras- length: 5–7 μm; pseudopore width: 7–12 μm.

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Figure 15. Mesonopora walteri sp. nov. (A–D) and Mesonopora aff. walteri sp. nov. (E, F) from the Callovian hardground of the Zalas Quarry, southern Poland.•A–general view of the sheet-like colony; B – autozooids; C, E – transversely elongate gonozooids with terminal ooeciopores (arrows); D, F – pseudopores. A–D – holotype, GIUS 8-3589-Z10/2; E, F – GIUS 8-3589-Z9/3.

Description. – Colony encrusting, irregularly discoidal, in quincunx, some closed by terminal diaphragms; unilamellar to multilamellar, rather thin, 5.5 to 7 mm in peristomes slightly raised, narrow (Fig. 14B). Pseudopores diameter. One small subcolony, which partly overgrows oval in shape, transversely elongate, closely spaced the parent colony, is present (Fig. 14A). (Fig. 14D). Autozooids cylindrical with slightly convex frontal Gonozooids numerous, situated mostly in the marginal walls crossed by prominent growth ridges 20 to 50 μm part of the colony, brood chamber roughly triangular or apart (Fig. 14B); autozooids longer in the central part of the transversely oval in shape (Fig. 14A, C). Ooeciopore oval, colony, more crowded and shorter around the margins. Ap- slightly longer then wide, smaller that an autozooidal aper- ertures oval in shape, slightly elongated distally, arranged ture (Fig. 14C).

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Remarks. – The specimens identified here as Hyporoso- Diagnosis. – Mesonopora with a sheet-like colony having pora cf. enstonensis have frontal walls covered by trans- elongate autozooids with widely spaced apertures; brood verse ridges, as in the type material of this species from the chamber broad, transversely elongated with slightly curved Bathonian Hampen Marly Beds of Oxfordshire, England distal margins; pseudopores semicircular. (see Pitt & Thomas 1969, p. 33, pl. 1, fig. 1, pl. 4, figs 2–3). However, the Zalas specimens have longer frontal walls Description. – Colony encrusting, sheet-like, berenici- and growth-ridges that are less distinct and regular than in form, multiserial, unilamellar, discoidal in outline H. enstonensis. Another species described from Poland (Fig. 15A). Autozooids elongate with slightly convex fron- with growth-ridges on the colony surface is Hyporosopora tal walls; zooidal boundaries well marked, shallowly groo- baltovensis Hara & Taylor, 1996 (see Hara & Taylor 1996, ved. Peristomes distinct, tapering distally. Apertures longi- pp. 95–98, figs 9–11, 13–18). In this species the ridges are tudinally elongated, some closed by pseudoporous discontinuous, irregularly spaced and 0.02–0.08 mm apart. diaphragms (Fig. 15B), distantly spaced and not aligned ra- Compared to H. cf. enstonensis, H. baltovensis has larger dially. Pseudopores slightly wider than longer, semicircu- autozooidal apertures and ooeciopores. lar in outline (Fig. 15D). Transverse ridges crossing the colony surface are found Gonozooids with flat proximal part and brood chamber in some other Jurassic species, such as Mesenteripora that is transversely elongated, six times wider than long, in- undulata (Michelin, 1845) (see Walter 1970), Plagioecia flated, the distal parts curving slightly towards the colony rugosa (d’Orbigny, 1853) and Hyporosopora portlandica interior, margins intended by numerous neighbouring (Gregory, 1896) (see Taylor 1981, p. 864, figs 3–6), which autozooids (Fig. 15C). Ooeciopore terminal, longitudinally are respectively Bathonian, Oxfordian and Tithonian in elongated, oval in shape, smaller than an autozooidal aper- age. The presence of ridges in Jurassic bryozoans was dis- ture, situated on a short ooeciostome. cussed by Hara & Taylor (1996) who concluded that the transversely ridged species seemed to be closely related Remarks. – Mesonopora concatenata (Reuss, 1867), a spe- and could be classified in the family Plagioeciidae (see cies known from the Upper Bathonian of Normandy and Hara & Taylor 1996). England (Walter 1970) and the Upper Bathonian–Lower Callovian of Balin, Poland (Taylor 2009), has similarly broad gonozooids but differs from the new species in its Genus Mesonopora Canu & Bassler, 1929 characteristically closely spaced apertures aligned in radial rows, smaller and more curved brood chambers, and trans- Type species. — Mesonopora concatenata (Reuss, 1867). versely elongated ooeciopore.

Mesonopora walteri sp. nov. Figure 15A–D Mesonopora aff. walteri sp. nov. Figure 15E, F Types. – Holotype: GIUS 8-3589-Z10/2 (Fig. 15A–D), paratype: GIUS 8-3589-M1. Material. – One colony encrusting a bivalve shell, GIUS 8-3589-Z9/3. Type locality. – Zalas Quarry, southern Poland. Measurements. – Autozooid frontal wall length: 310–520 μm; Type horizon. – Callovian hardground, Middle Jurassic. autozooid frontal wall width: 100–140 μm; longitudinal apertural diameter: 90–125 μm; transverse apertural dia- Etymology. – In honour of Bernard Walter, French resear- meter: 62–87 μm; gonozooid total length: 650 μm; brood cher on Mesozoic bryozoans. chamber length: 400 μm; gonozooid width: 1850 μm; ooeciopore length: 50 μm; ooeciopore width: 50 μm; pseudo- Material. – Two specimens encrusting bivalve shells: pore length: 5 μm; pseudopore width: 10 μm. GIUS 8-3589-Z10/2 and GIUS 8-3589-M1. Remarks. – This taxon is similar to Mesonopora walteri Measurements. – Autozooid frontal wall length: 490–690 μm; sp. nov. described above but differs in having smaller auto- autozooid frontal wall width: 160–210 μm; longitudinal zooids and transversely elliptical pseudopores that are two apertural diameter: 110–125 μm; transverse apertural dia- times wider than long (Fig. 15F). Assuming that pseudopore meter: 75–87 μm; gonozooid total length: 1090 μm; brood characteristics are species specific (e.g., Taylor & Zatoń chamber length: 580 μm; gonozooid width: 3850 μm; ooe- 2008, Zatoń & Taylor 2009), then this colony could repre- ciopore length: 90 μm; ooeciopore width: 60 μm; pseudo- sent a distinct species of Mesonopora but additional speci- pore length: 6–9 μm; pseudopore width: 7–10 μm. mens are needed to warrant introduction of a new species.

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Figure 16. Theonoa minuta (Reuss, 1867) from the Callovian hardground of the Zalas Quarry, southern Poland, GIUS 8-3589-M15b. • A – fasciculate colony view; B – fascicles arrangement; C – autozooids; D – pseudopores.

Family Theonoidae Busk, 1859 Description. – Colonies encrusting, multiserial, unilamellar, circular in outline (Fig. 16A), small (3.6 mm in diame- Genus Theonoa Lamouroux, 1821 ter), fasciculate. Fascicles uniserial in early astogeny, becoming multiserial; new fascicles arising through Type species. – Theonoa chlatrata Lamouroux, 1821 by bifurcation or intercalation (Fig. 16B). Autozooids small monotypy. with rounded-quadrate apertures 80–120 μm in diameter (Fig. 16C). Gonozooids not observed. Pseudopores badly Theonoa minuta (Reuss, 1867) preserved but apparently subcircular in outline (Fig. 16D), Figure 16A–D about 8 μm in diameter.

1867 Pavotubigera minuta, Reuss, p. 3, pl. 2, fig. 8. Remarks. – The species was revised by Taylor (2009) 1970 Theonoa chlatrata Lamouroux. – Walter, p. 190 (par- based on the original material of Reuss (1867) from Balin. tim), non pl. 19, figs 8, 9, 11, non pl. 20, fig. 1 [= Theo- The colonies from the Callovian of Zalas described here, noa chlatrata Lamouroux, 1821]. although infertile, match the Balin type well. The Bathon- 2009 Theonoa minuta (Reuss). – Taylor, p. 44, fig. 12A–F. ian species Theonoa chlatrata Lamouroux, 1821, origi- nally described from Normandy (Walter 1970), differs Material. – Three neighbouring colonies encrusting a bi- from T. minuta in developing larger colonies, autozooids valve shell, GIUS 8-3589-M15b. and gonozooids (see also Taylor 2009).

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with transverse growth bands (Fig. 18B, C); pseudopores seemingly lacking (Fig. 18C, D). Gonozooids not observed.

Remarks. – In the absence of gonozooids, this form is assigned to the informal genus “Berenicea” following the recommendation of Taylor & Sequeiros (1982). A re- markable feature of the two colonies from Zalas is their apparent lack of pseudopores. Other cyclostome species from Zalas show clear pseudopores, even when the sur- faces of the colonies are only moderately well-preserved. One of the two specimens (GIUS 8-3589-Z11/3) exhi- bits particularly good preservation of the frontal walls, 200 µm yet no pseudopores are visible. Cyclostome pseudopores can be sparse and are occasionally partly or completely Figure 17. Undeterminable entalophorid base, GIUS 8-3589-Z9/1. occluded, as in Stomatopora cf. dichotomoides (d’Orbigny, 1850) described above, but in such instan- ces they tend to be fully open and clearly visible at the Family Entalophoridae Reuss, 1869 growing edge, which is not the case in the Zalas “Bereni- cea”. Two possibilities can be suggested: either the Entalophoridae gen. et. sp. indet. pseudopores in the Zalas forms are very sparsely distri- Figure 17 buted and have been overlooked, or they are absent alto- gether. Material. – Six bases attached to bivalve shells, of which To our knowledge all post-Palaeozoic cyclostomes two have been SEMed: GIUS 8-3589-Z9/1-2. with exterior walls possess pseudopores. There are, however, Palaeozoic cyclostomes that lack pseudopores, Remarks. – The present material consists of poorly preser- including the Ordovician-Devonian genus Sagenella ved bases of erect cyclostomes that are indeterminate at ge- (Suborder Paleotubuliporina), which strongly resembles nus level. The absence of an axial lumen in the erect bran- some post-Palaeozoic Berenicea-like cyclostomes ex- ches indicates that the specimens do not belong to the cept for the absence of pseudopores and gonozooids. genus Entalophora itself (see Walter 1970, Taylor & The best-known species, Sagenella consimilis (Lons- McKinney 2006, Zatoń & Taylor, 2010). dale), which occurs in the Wenlock of Britain and Gotland, Sweden, is superficially very similar to the Zalas forms of “Berenicea”, although each zooid has a Family Incertae sedis pair of minute, raised pores along the midline (see Brood 1975). It would be premature to assert a close affinity be- “Berenicea” sp. tween the Zalas taxa and Palaeozoic Sagenella but fur- Figure 18A–D ther research is warranted to test the possibility that paleotubuliporine cyclostomes lacking pseudopores sur- Material. – Two colonies encrusting bivalve shells: GIUS vived into the Jurassic. 8-3589-Z11/3, GIUS 8-3589-Z7/1.

Description. – Colony encrusting, multiserial, sheet-like, Suborder Cerioporina Hagenov, 1851 unilamellar, subcircular in outline (Fig. 18A), about Family Cavidae d’Orbigny, 1854 4.5 mm in maximum diameter, thin with growing edge nor- mally exposing only one generation of zooids, basal lamina Genus Ceriocava d’Orbigny, 1854 not extending significantly beyond budding zone, pustules visible on basal walls of some new buds. Early astogenetic Ceriocava sp. stages obscured by sediment. Figure 19A, B Autozooids slender (Fig. 18B), up to 800 μm long and attaining 140 μm in maximum width close to the short pre- Material. – Several variably preserved colonies, including served peristome. Apertures arranged quincuncially, longi- the figured specimen GIUS 8-3589-C/W-230/3. tudinally elongate, up to 100 μm long by 70 μm wide, some closed by terminal diaphragms (Fig. 18B). Frontal wall Description. – Colonies dome-shaped, free-walled with

858 Micha³ Zatoñ et al. Callovian (Middle Jurassic) cyclostome bryozoans from southern Poland

200 µm A 100 µm B

10 µm C 20 µm D

Figure 18. “Berenicea” sp., a bereniciform cyclostome of uncertain affinity from the Callovian hardground of the Zalas Quarry, southern Poland, GIUS 8-3589-Z11/3. • A – sheet-like colony view; B – autozooids; C, D – autozooidal frontal walls apparently devoid of pseudopores. autozooids openings over entire upper surface (Fig. 19A). Family Heteroporidae Waters, 1880 One colony preserves the ancestrula with a smooth, circular, bulbous protoecium 270 μm in diameter (Fig. 19B), se- Genus Ripisoecia Canu & Bassler, 1922 parated from the rest of the colony by a distinct constriction. Autozooids with rounded polygonal apertures and Ripisoecia conifera (Lamouroux, 1821) thick interzooidal walls giving the colony a characteristic Figure 19C, D honeycomb-like pattern (Fig. 19A). Gonozooids not observed. 1821 Millepora conifera, Lamouroux, p. 87, pl. 83, figs 6–7. Remarks. – Colony shape and autozooidal characteristics 1867 Heteropora conifera Lamouroux. – Reuss, p. 12 (par- resemble other representatives of the genus (e.g., Walter tim), pl. 1, fig. 12, pl. 2, fig. 1 only. 1970; Taylor 2009; Zatoń & Taylor 2009, 2010). However, 1970 Ripisoecia conifera (Lamouroux). – Walter, p. 160, the lack of gonozooids precludes species identification. pl. 16, figs 6–12, pl. 17, figs 1, 2. Thus, it cannot be determined whether the Zalas Callovian 2009 Ripisoecia conifera (Lamouroux). – Taylor, p. 48, specimens represent the well-known species Ceriocava co- figs 13B, 14D–F. rymbosa (Lamouroux, 1821), or are a different species, as is also the case for Ceriocava specimens in the Upper Bajo- Material. – One colony encrusting a bivalve shell, GIUS cian–Upper Bathonian of Poland (Zatoń & Taylor 2009, 8-3589-Z7/2. 2010). It is worth noting that the Upper Bathonian–Lower Callovian specimens of Ceriocava specimens from the Description. – Colony globular, fungiform (Fig. 19C), nearby Balin locality (see Taylor 2009) lack gonozooids, 4 mm in diameter, attached to the substrate by a short stalk. making their assignment to Ceriocava corymbosa also un- Apertures rounded polygonal in outline, separated by thick certain. interzooidal walls; autozooidal apertures up to 220 μm in

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100 µm A 100 µm B

200 µm C 100 µm D

Figure 19. Ceriocava sp. (A, B) and Ripisoecia conifera (Lamouroux, 1821) (C, D) from the Callovian hardground of the Zalas Quarry, southern Po- land.•A–juvenile colony; B – protoecium; C – dome-shaped colony; D – autozooidal apertures and smaller kenozooidal apertures (e.g., arrow). A, B – GIUS 8-3589-C/W-230/3; C, D – GIUS 8-3589-Z7/2. diameter interspersed with smaller kenozooidal apertures strate space they occupy (see Jackson 1979, Taylor 1999) up to 80 μm (Fig. 19D). and consist of such bereniciform genera as Microeciella, Hyporosopora, Mesonopora and Reptomultisparsa, plus ge- Remarks. – This species was recorded from the classic lo- nerically indeterminate forms assignable to the form-genus cality at Balin, southern Poland by Reuss (1867), revised “Berenicea”. Oligoserial branching colonies, some of which by Taylor (2009). It differs from the related Ceriocava in can be assigned to Oncousoecia, have an intermediate mor- its globular to fungiform colonies and dimorphic apertures. phological strategy. Also present at Zalas are the massive, Gonozooids have not been seen in either the Balin or Zalas often multilayered, dome-shaped and fungiform genera of colonies. Ceriocava and Ripisoecia, respectively. The fasciculate genus Theonoa and indeterminate “entalophorids” are minor constituents in the Zalas assemblage. Discussion and conclusions Bereniciform cyclostomes, although often numerous on a given hard substrate, are mainly represented by infer- As in other Jurassic cyclostome bryozoan assemblages, the tile colonies that are indeterminate at genus-level. For ex- fauna from the Callovian of Zalas is represented by three do- ample, of the 357 bereniciform colonies from the Callovian minant encrusting colony-forms: uniserial branching run- of Zalas examined by Zatoń et al. (2011), only 37 (10%) ners, oligoserial branching ribbons, and multiserial sheets. were fertile. This pattern is repeated in other Jurassic as- The runner-like colonies are fugitive strategists, which are semblages (e.g., Zatoń & Taylor 2009, 2010; Hara 2007; well-adapted to locating spatial refuges on the substrate (see Hara & Taylor 2009), impacting analyses of assemblage Buss 1979, Jackson 1979, Taylor 1999). They are mainly re- diversity. presented by species of Stomatopora. In contrast, the The Callovian cyclostome assemblage described here sheet-like colonies are better adapted to retaining the sub- from Zalas is represented by at least twenty-two taxa. Co-

860 Micha³ Zatoñ et al. Callovian (Middle Jurassic) cyclostome bryozoans from southern Poland incidentally, the cyclostome fauna from the condensed Up- 2001; see also Sogot et al. 2013), the clear predominance per Bathonian–Lower Callovian Oolite at the nearby local- of bereniciform colonies over the refuge-seeking runners ity of Balin contains a very similar number (23) of species (see Zatoń et al. 2011, table 1) may be indicative of a rather (Taylor 2009). In the context of the Jurassic, Zalas and eutrophic conditions prevailing in the basin during the Balin are diverse: few other Jurassic bryozoan faunas con- times of the hardground formation. Similar conclusions, as tain more than 20 species (Taylor & Wilson 1999). An- based on the presence of an abundant and diverse other diverse Callovian cyclostome bryozoan fauna comes polychaete tubes, oysters and bryozoans, were reached by from the Moscow region of Russia where 20 species were Zatoń et al. (2011) on the basis of criteria worked out ear- described by Gerasimov (1955) and are currently under re- lier in a Recent sclerobiont biota by Lescinsky et al. vision by Viskova (e.g., Viskova 2006, 2007, 2008, 2009). (2002). The Balin Oolite, although containing some genera The strikingly patchy distribution of Jurassic bryozoan (Multisparsa, Idmonea and Mesenteripora) not present in faunas in time and space (Taylor & Ernst 2008) makes de- the Zalas assemblage also possesses a few species in com- scriptions of faunas such as from Zalas especially impor- mon, such as Hyporosopora tenera, Ripisoecia conifera tant for improving our knowledge of evolutionary and di- and Theonoa minuta. The Zalas stomatoporids, although versity patterns during this unique time when cyclostome only identifiable with uncertainty, are also very similar to bryozoans achieved dominance before the advent of the those from Balin. Erect “entalophorids” that are unknown cheilostomes. The assemblages from Zalas and Balin con- from the Balin assemblage occur in the Zalas bryozoan as- tain the highest diversity of Callovian cyclostome bryo- semblage (Zatoń et al. 2011). Hyporosopora tenera and zoans known to date and thus serve as a reference for diver- other species similar to those previously described from sity comparisons with subsequent work on Callovian somewhat older Bathonian strata of the Polish Jura (see assemblages. Zatoń & Taylor 2009, 2010), such as Microeciella aff. annae, M. cf. maleckii and Reptomultisparsa aff. kawodrzanensis, occur in the Callovian assemblage of Zalas. Acknowledgments Differences in species composition between Balin and Zalas may reflect different environmental settings (shal- This research was possible due to the financial support of the Fac- lower water in Balin than Zalas). However, it must also be ulty of Earth Sciences to MZ. Laís Ramalho and Kamil Zágoršek stressed that a large number of cyclostome colonies from commented on the early version of the manuscript, providing nu- Zalas are infertile and thus many other genera and species merous remarks and corrections that are greatly appreciated. We å may have existed that have gone unrecognized, causing are grateful to Roger J. Cuffey and Eckart H kansson, the journal reviewers, for further useful remarks and comments that im- true diversity to be underestimated. proved the present article. We also thank Mrs Barbara Both environmental (unchanging salinity, reduced al- Ruszkiewicz for taking the photographs of the bryozoan-en- gal cover) and depositional (reduced sedimentation rate in crusted shells. a deeper setting) conditions were important for the establishment of diverse sclerobiont communities in the Cal- lovian of the Zalas area (Zatoń et al. 2011). The prolonged References interval of non-deposition during hardground formation (e.g., Giżejewska & Wieczorek 1976, Matyja 2006, BASSLER, R.S. 1935. Bryozoa. Generum et Genotyporum. Index Dembicz & Praszkier 2007) was one of the most important et Bibliographica, 1–229. In QUENSTEDT, W. (ed.) Fossilium factors, resulting in the presence of abundant secondary Catalogus I. Animalia. Vol. Part 67. W. Junk, s’Gravenhage. hard substrates for colonization. A problem yet to be re- BRONN, H.G. 1825. System der urweltlichten Pflanzenthiere solved is how so many morphologically similar durch Diagnose, Analyse und Abbildung der Geschlechter bereniciform taxa were able to coexist in Jurassic bryozoan erläutert. 47 pp. Mohr, Heidelberg. assemblages, including that described here from Zalas. BROOD, K. 1975. Cyclostomatous Bryozoa from the Silurian of These forms exhibit a common strategy for utilizing sub- Gotland. Stockholm Contributions in Geology 28, 45–119. BUSK, G. 1852. An account of the Polyzoa and Sertularian Zoo- strate space (cf. the refuge-seeking strategy of Stoma- phytes collected in the voyage of the Rattlesnake on the coast topora; see Taylor 1979, 1999), and have similar life-histo- of Australia and the Louisiade Archipelago, 343–402. In ries (see McKinney & Taylor 1997). In addition the sizes of MACGILLIVRAY, J. (ed.) Narrative of the Voyage of H.M.S. Rat- the zooids spans too narrow a range to support significant tlesnake, commanded by the late Captain Owen Stanley, dur- niche partitioning by food particle size. The functional sig- ing the years 1846–1850, 1. Boone, London. nificance of differences in pseudopore morphology be- BUSK, G. 1859. A monograph of the fossil Polyzoa of the Crag. tween cyclostome species has yet to be understood. 136 pp. Palaeontographical Monograph, London. As runners such as Stomatopora are considered to be DOI 10.5962/bhl.title.2037 well-adapted to low nutrient environments (Okamura et al. BUSS, L.W. 1979. 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